Sauropod dinosaur tracks from the Purbeck Group (Early Cretaceous) of Spyway Quarry, Dorset, UK.

Dinosaur tracks have a long history of discovery and study in the UK, but track sites for sauropodomorph dinosaurs-the group that included the giant, graviportal herbivorous sauropods-are comparatively rare. Here, we provide a description of a sauropod dinosaur track site at Spyway Quarry in Dorset, southern England. The tracks at Spyway were discovered in the late 1990s and occur in the Stair Hole Member of the Durlston Formation in the Purbeck Limestone Group, of earliest Cretaceous age. More than 130 individual tracks of large sauropod dinosaurs are present at the site, but they are generally poorly preserved and do not form clear trackways, although it is likely that they represent multiple individuals. They provide further evidence for sauropods living in or passing through coastal lagoonal environments. Although poorly preserved, Spyway represents the largest in situ dinosaur track site currently accessible within the Purbeck Group, with considerable potential for further discoveries through ongoing quarrying in the surrounding area.

A new semi-fossorial thescelosaurine dinosaur from the Cenomanian-age Mussentuchit Member of the Cedar Mountain Formation, Utah.

Thescelosaurines are a group of early diverging, ornithischian dinosaurs notable for their conservative bauplans and mosaic of primitive features. Although abundant within the latest Cretaceous ecosystems of North America, their record is poor to absent in earlier assemblages, leaving a large gap in our understanding of their evolution, origins, and ecological roles. Here we report a new small bodied thescelosaurine-Fona herzogae gen. et sp. nov.-from the Mussentuchit Member of the Cedar Mountain Formation, Utah, USA. Fona herzogae is represented by multiple individuals, representing one of the most comprehensive skeletal assemblages of a small bodied, early diverging ornithischian described from North America to date. Phylogenetic analysis recovers Fona as the earliest member of Thescelosaurinae, minimally containing Oryctodromeus, and all three species of Thescelosaurus, revealing the clade was well-established in North America by as early as the Cenomanian, and distinct from, yet continental cohabitants with, their sister clade, Orodrominae. To date, orodromines and thescelosaurines have not been found together within a single North American ecosystem, suggesting different habitat preferences or competitive exclusion. Osteological observations reveal extensive intraspecific variation across cranial and postcranial elements, and a number of anatomical similarities with Oryctodromeus, suggesting a shared semi-fossorial lifestyle.

A new stegosaur (Dinosauria: Ornithischia) from the Middle Jurassic of Gansu Province, China.

Stegosaurs are a minor but iconic clade of ornithischian dinosaurs, yet due to a poor fossil record, their early evolution is poorly understood. Here, we describe a new stegosaur, Baiyinosaurus baojiensis, gen. et sp. nov. from the Middle Jurassic Wangjiashan Formation of the Pingchuan District, Baiyin City, Gansu Province, China. The frontal of Baiyinosaurus possesses a unique characteristic among Stegosauria: it is wider than long and contributes to both the medial and anterior margins of the supratemporal fenestra. The character combinations of dorsal vertebrae of Baiyinosaurus are also different to other stegosaurs: its neural arches are not greatly elongated, its parapophyses are well developed, and its neural spines are axially expanded in lateral. The features of the frontal and vertebrae of Baiyinosaurus are reminiscent of basally branching thyreophorans, indicating that Baiyinosaurus is transitional in morphology between early thyreophorans and early-diverging stegosaurs. Systematic analysis shows that Baiyinosaurus is an early-diverging stegosaur.

Exceptional Early Jurassic fossils with leathery eggs shed light on dinosaur reproductive biology.

Our understanding of pre-Cretaceous dinosaur reproduction is hindered by a scarcity of evidence within fossil records. Here we report three adult skeletons and five clutches of embryo-containing eggs of a new sauropodomorph from the Lower Jurassic of southwestern China, displaying several significant reproductive features that are either unknown or unlike other early-diverging sauropodomorphs, such as relatively large eggs with a relatively thick calcareous shell formed by prominent mammillary cones, synchronous hatching and a transitional prehatching posture between the crocodilians and living birds. Most significantly, these Early Jurassic fossils provide strong evidence for the earliest known leathery eggs. Our comprehensive quantitative analyses demonstrate that the first dinosaur eggs were probably leathery, elliptical and relatively small, but with relatively long eggshell units, and that along the line to living birds, the most significant change in reptilian egg morphology occurred early in theropod evolution rather than near the origin of Aves.

An unusually robust specimen attributed to Buriolestes schultzi (Dinosauria: Sauropodomorpha) from the Late Triassic of southern Brazil.

Buriolestes schultzi is a small sauropodomorph dinosaur from Carnian beds (ca., 233 Ma) of southern Brazil. It is one of the earliest members of that lineage and is a key taxon to investigate the initial evolution of Sauropodomorpha. Here, we attribute a new specimen to B. schultzi from Late Triassic of southern Brazil, which represents the first occurrence of the taxon outside the type locality. The new specimen comprises a disarticulated and partial skeleton, including cranial and postcranial elements. It is tentatively regarded as an additional specimen of B. schultzi according to a unique combination of traits (including autapomorphies). Conversely, the new specimen is stouter than the other specimens of B. schultzi, as shown by femoral Robustness Index. Based on femoral circumference, the estimated body mass of the new specimen is approximately 15 kg, which is far higher than the previous estimations for other specimens of B. schultzi (i.e., approximately 7 kg). In fact, the new specimen and some specimens of Eoraptor lunensis and Saturnalia tupiniquim were found to be significantly stouter than coeval sauropodomorphs. Therefore, instead of all being constructed as gracile, the earliest sauropodomorphs experienced an unappreciated intraspecific variation in robustness. This is interesting because more precise data on species body mass are crucial in order to better understand the complex terrestrial ecosystems in which dinosaurs originated.

New Middle to ?Late Jurassic dinosaur tracksites in the Central High Atlas Mountains, Morocco.

Besides bones, fossil tracks and trackways are important sources of knowledge about dinosaur palaeobiology. Here, we report three new tracksites from two different synclines in the Imilchil area, Central High Atlas, Morocco. The tracks and trackways are preserved in fluvial deposits in different levels of the Isli Formation (Early Bathonian-?Upper Jurassic), and contain impressions made by sauropods, theropods and ornithopods, as well as tracks that might represent bird-like non-avian theropod dinosaurs. In addition to traditional field measurements, three-dimensional digital models of the track sites were created using photogrammetry. These new tracksites add to the rich faunal ichnoassemblage already recorded from the High Atlas Mountains and North Africa, which is considerably richer than the contemporaneous body fossil record, and also provide new data on dinosaurs-substrate interactions.

Osteohistology reveals the smallest adult Jurassic sauropodomorph.

The earliest sauropodomorphs were small omnivores (less than 10 kg) that first appeared in the Carnian. By the Hettangian, early branching sauropodomorphs (EBSMs) were globally distributed, had variable postures, and some attained large body masses (greater than 10 tonnes). Small-bodied EBSMs like Massospondylus carinatus (less than 550 kg) persist at least until the Pliensbachian at nearly all dinosaur-bearing localities worldwide but are comparatively low in alpha diversity. One potential reason for this is competition with other similarly sized contemporary amniotes, including Triassic gomphodont cynodonts, Jurassic early branching ornithischians, herbivorous theropods and potentially early crocodylomorphs. Today's herbivorous mammals show a range of body size classes (less than 10 g to 7 tonnes), with multiple species of small herbivorous mammals (less than 100 kg) frequently co-occurring. Comparatively, our understanding of the phylogenetic distribution of body mass in Early Jurassic strata, and its explanatory power for the lower thresholds of body mass in EBSMs, needs more data. We osteohistologically sectioned a small humerus, BP/1/4732, from the upper Elliot Formation of South Africa. Its comparative morphology and osteohistology show that it represents a skeletally mature individual of a new sauropodomorph taxon with a body mass of approx. 75.35 kg. This makes it one of the smallest known sauropodomorph taxa, and the smallest ever reported from a Jurassic stratum.

Dinosaur trackways from the Upper Cretaceous Nichkesai Formation near Mayluu Suu City, Southern Tien Shan Mountains, north-western Kyrgyzstan.

Trackways provide essential data on the biogeographic distribution, locomotion and behaviour of dinosaurs. Cretaceous dinosaur trackways are abundant in the Americas, Europe, North Africa and East Asia, but are less well documented in Central Asia despite extensive exposure of Cretaceous terrestrial sedimentary rocks in the region. Here we report the presence of bipedal, tridactyl dinosaur trackways near the city of Mayluu Suu, Jalal Abad Oblast, north-western Kyrgyzstan, the first discovery of dinosaur trace fossils within the country. The trackways are situated on a steep slope uncovered by a landslide around the year 2000 in a highly landslide-affected area. Photogrammetry is used to digitally analyse and conserve the trace fossils. We infer a shoreface setting for the trackways based on the locality sedimentology, discuss the identity of the track makers and highlight the potential for future trackway discovery in the area. This discovery contributes vital data to an otherwise sparse record on the spatio-temporal distribution of dinosaurs in Kyrgyzstan, and to the dinosaur trackway record of Central Asia.

Comparative microstructural study on the teeth of Mesozoic birds and non-avian dinosaurs.

Although it is commonly considered that, in birds, there is a trend towards reduced dentition, teeth persisted in birds for 90 Ma and numerous macroscopic morphologies are observed. However, the extent to which the microstructure of bird teeth differs from other lineages is poorly understood. To explore the microstructural differences of the teeth of birds in comparison with closely related non-avialan dinosaurs, the enamel and dentine-related features were evaluated in four Mesozoic paravian species from the Yanliao and Jehol biotas. Different patterns of dentinal tubular tissues with mineralized extensions of the odontoblast processes were revealed through the examination of histological sectioning under electron microscopy. Secondary modification of the tubular structures, forming reactive sclerotic dentin of Longipteryx, and the mineralization of peritubular dentin of Sapeornis were observed in the mantle dentin region. The new observed features combined with other dentinal-associated ultrastructure suggest that the developmental mechanisms controlling dentin formation are quite plastic, permitting the evolution of unique morphologies associated with specialized feeding behaviours in the toothed birds. Proportionally greater functional stress placed on the stem bird teeth may have induced reactive dentin mineralization, which was observed more often within tubules of these taxa. This suggests modifications to the dentin to counteract potential failure.

The selfish preen: absence of allopreening in Palaeognathae and its socio-cognitive implications.

Preening behaviours are widespread in extant birds. While most birds appear to autopreen (self-directed preening), allopreening (preening directed at conspecifics) seems to have emerged only in certain species, but across many families. Allopreening has been hypothesised to reinforce mutual relationships and cooperation between individuals, and to underpin various socio-cognitive abilities. Palaeognathae is a bird group exhibiting neurocognitively plesiomorphic traits compared to other birds. They share many features with non-avian paravian dinosaurs and are thus important for the study of cognitive evolution in birds. Despite this, and the important correlation of allopreening with many complicated social behaviours, allopreening has not been systematically studied in Palaeognathae. Therefore, we examined the preening behaviours in four species of palaeognaths: common ostriches (Struthio camelus), greater rheas (Rhea americana), emus (Dromaius novaehollandiae), and elegant crested tinamous (Eudromia elegans). We compared findings with common ravens (Corvus corax), a neognath species known for its allopreening and complex social cognition. We found autopreening, but no allopreening, in the palaeognath species, while both autopreening and allopreening was found in common ravens. The absence of allopreening in Palaeognathae suggests an emergence of this behaviour within Neognathae. We contextualise our results in relation to the socio-cognitive underpinnings of allopreening and its implications for the understanding of the evolution of socio-cognitive abilities in non-avian paravian dinosaurs and early birds.

How smart dinosaurs?

This commentary discusses the main points made in Reiner's article on the prospect that some theropod dinosaurs could have given rise to a lineage that achieved a human level of intelligence, and those made in Herculano-Houzel's article on the potentially monkey-like numbers of neurons in the pallium of large theropods, and the implications of this for their intelligence.

Could theropod dinosaurs have evolved to a human level of intelligence?

Noting that some theropod dinosaurs had large brains, large grasping hands, and likely binocular vision, paleontologist Dale Russell suggested that a branch of these dinosaurs might have evolved to a human intelligence level, had dinosaurs not become extinct. I offer reasons why the likely pallial organization in dinosaurs would have made this improbable, based on four assumptions. First, it is assumed that achieving human intelligence requires evolving an equivalent of the about 200 functionally specialized cortical areas characteristic of humans. Second, it is assumed that dinosaurs had an avian nuclear type of pallial organization, in contrast to the mammalian cortical organization. Third, it is assumed that the interactions between the different neuron types making up an information processing unit within pallium are critical to its role in analyzing information. Finally, it is assumed that increasing axonal length between the neuron sets carrying out this operation impairs its efficacy. Based on these assumptions, I present two main reasons why dinosaur pallium might have been unable to add the equivalent of 200 efficiently functioning cortical areas. First, a nuclear pattern of pallial organization would require increasing distances between the neuron groups corresponding to the separate layers of any given mammalian cortical area, as more sets of nuclei equivalent to a cortical area are interposed between the existing sets, increasing axon length and thereby impairing processing efficiency. Second, because of its nuclear organization, dinosaur pallium could not reduce axon length by folding to bring adjacent areas closer together, as occurs in cerebral cortex.

A potentially fatal cranial pathology in a specimen of Tarchia.

Skulls of the Mongolian ankylosaurids Shamosaurus, Tarchia, and Saichania were scanned for information about their internal anatomy. Computed tomography (CT) imaging of the Tarchia skull revealed substantial internal anatomical differences from known Campanian North American taxa, particularly in the morphology of the airway. In addition, unexpected anomalies were detected within the airway and sinuses. The anomalies include multiple bilaterally distributed, variably sized hyperdense (mineralized) concretions within the airway and sinuses, the largest of which, positioned in the right nasal cavity medial to the supraorbitals, has an asymmetric ovoid shape that tapers caudally and which is partially encased within a hemispherical trabeculated osseous proliferation (sinus exostosis). Immediately adjacent to the exostosis is a subcircular transosseous defect in the prefrontal region of the skull roof that is partially filled with trabeculated ossified material with similar architectural features as the larger exostosis. Irregularities along the internal and external surfaces of the cranial vault may be associated. The radiologic features of the hemicircumferential exostosis suggest a chronic reactive osteoproliferation, possibly in response to an ongoing inflammatory reaction to primary sinus infection or, in combination with the unilateral transosseous defect, traumatically introduced infection with potentially fatal consequences. This report underscores the value of CT scanning of fossil vertebrate specimens, which in this case revealed large internal lesions of the skull that, at the time the scan was performed, were otherwise indiscernible.

Symbiosis between Cretaceous dinosaurs and feather-feeding beetles.

Extant terrestrial vertebrates, including birds, have a panoply of symbiotic relationships with many insects and arachnids, such as parasitism or mutualism. Yet, identifying arthropod-vertebrate symbioses in the fossil record has been based largely on indirect evidence; findings of direct association between arthropod guests and dinosaur host remains are exceedingly scarce. Here, we present direct and indirect evidence demonstrating that beetle larvae fed on feathers from an undetermined theropod host (avian or nonavian) 105 million y ago. An exceptional amber assemblage is reported of larval molts (exuviae) intimately associated with plumulaceous feather and other remains, as well as three additional amber pieces preserving isolated conspecific exuviae. Samples were found in the roughly coeval Spanish amber deposits of El Soplao, San Just, and Peñacerrada I. Integration of the morphological, systematic, and taphonomic data shows that the beetle larval exuviae, belonging to three developmental stages, are most consistent with skin/hide beetles (family Dermestidae), an ecologically important group with extant keratophagous species that commonly inhabit bird and mammal nests. These findings show that a symbiotic relationship involving keratophagy comparable to that of beetles and birds in current ecosystems existed between their Early Cretaceous relatives.

Evidence for heterothermic endothermy and reptile-like eggshell mineralization in Troodon, a non-avian maniraptoran theropod.

The dinosaur-bird transition involved several anatomical, biomechanical, and physiological modifications of the theropod bauplan. Non-avian maniraptoran theropods, such as Troodon, are key to better understand changes in thermophysiology and reproduction occurring during this transition. Here, we applied dual clumped isotope (Δ47 and Δ48) thermometry, a technique that resolves mineralization temperature and other nonthermal information recorded in carbonates, to eggshells from Troodon, modern reptiles, and modern birds. Troodon eggshells show variable temperatures, namely 42 and 29 ± 2 °C, supporting the hypothesis of an endothermic thermophysiology with a heterothermic strategy for this extinct taxon. Dual clumped isotope data also reveal physiological differences in the reproductive systems between Troodon, reptiles, and birds. Troodon and modern reptiles mineralize their eggshells indistinguishable from dual clumped isotope equilibrium, while birds precipitate eggshells characterized by a positive disequilibrium offset in Δ48. Analyses of inorganic calcites suggest that the observed disequilibrium pattern in birds is linked to an amorphous calcium carbonate (ACC) precursor, a carbonate phase known to accelerate eggshell formation in birds. Lack of disequilibrium patterns in reptile and Troodon eggshells implies these vertebrates had not acquired the fast, ACC-based eggshell calcification process characteristic of birds. Observation that Troodon retained a slow reptile-like calcification suggests that it possessed two functional ovaries and was limited in the number of eggs it could produce; thus its large clutches would have been laid by several females. Dual clumped isotope analysis of eggshells of extinct vertebrates sheds light on physiological information otherwise inaccessible in the fossil record.

How Triceratops got its face: An update on the functional evolution of the ceratopsian head.

Ceratopsian dinosaurs arguably show some of the most extravagant external cranial morphology across all Dinosauria. For over a century, ceratopsian dinosaurs have inspired a multitude of cranial functional studies as more discoveries continued to depict a larger picture of the enormous diversity of these animals. The iconic horns and bony frills in many ceratopsians portray a plethora of shapes, sizes, and arrangements across taxa, and their overall feeding apparatus show the development of unique specializations previously unseen in large herbivores. Here, I give a brief updated review of the many functional studies investigating different aspects of the ceratopsian head. The functional role of the horns and bony frills is explored, with an overview of studies investigating their potential for weaponization or defense in either intraspecific or anti-predatory combat, among other things. A review of studies pertaining to the ceratopsian feeding apparatus is also presented here, with analyses of studies exploring their beak and snout morphology, dentition and tooth wear, cranial musculature with associated skull anatomy, and feeding biomechanics.

Investigating the quadrupedal abilities of Scutellosaurus lawleri and its implications for locomotor behavior evolution among dinosaurs.

A reversion to secondary quadrupedality is exceptionally rare in nature, yet the convergent re-evolution of this locomotor style occurred at least four separate times within Dinosauria. Facultative quadrupedality, an intermediate state between obligate bipedality and obligate quadrupedality, may have been an important transitional step in this locomotor shift, and is proposed for a range of basal ornithischians and sauropodomorphs. Advances in virtual biomechanical modeling and simulation have allowed for the investigation of limb anatomy and function in a range of extinct dinosaurian species, yet this technique has not been widely applied to explore facultatively quadrupedal gait generation. This study places its focus on Scutellosaurus, a basal thyreophoran that has previously been described as both an obligate biped and a facultative quadruped. The functional anatomy of the musculoskeletal system (myology, mass properties, and joint ranges of motion) has been reconstructed using extant phylogenetic bracketing and comparative anatomical datasets. This information was used to create a multi-body dynamic locomotor simulation that demonstrates that whil quadrupedal gaits were physically possible, they did not outperform bipedal gaits is any tested metric. Scutellosaurus cannot therefore be described as an obligate biped, but we would predict its use of quadrupedality would be very rare, and perhaps restricted to specific activities such as foraging. This finding suggests that basal thyreophorans are still overwhelmingly bipedal but is perhaps indicative of an adaptive pathway for later evolution of quadrupedality.

Theropod dinosaurs had primate-like numbers of telencephalic neurons.

Understanding the neuronal composition of the brains of dinosaurs and other fossil amniotes would offer fundamental insight into their behavioral and cognitive capabilities, but brain tissue is only rarely fossilized. However, when the bony brain case is preserved, the volume and therefore mass of the brain can be estimated with computer tomography; and if the scaling relationship between brain mass and numbers of neurons for the clade is known, that relationship can be applied to estimate the neuronal composition of the brain. Using a recently published database of numbers of neurons in the telencephalon of extant sauropsids (birds, squamates, and testudines), here I show that the neuronal scaling rules that apply to these animals can be used to infer the numbers of neurons that composed the telencephalon of dinosaur, pterosaur, and other fossil sauropsid species. The key to inferring numbers of telencephalic neurons in these species is first using the relationship between their estimated brain and body mass to determine whether bird-like (endothermic) or squamate-like (ectothermic) rules apply to each fossil sauropsid species. This procedure shows that the notion of "mesothermy" in dinosaurs is an artifact due to the mixing of animals with bird-like and squamate-like scaling, and indicates that theropods such as Tyrannosaurus and Allosaurus were endotherms with baboon- and monkey-like numbers of telencephalic neurons, respectively, which would make these animals not only giant but also long-lived and endowed with flexible cognition, and thus even more magnificent predators than previously thought.

Iyuku raathi, a new iguanodontian dinosaur from the Early Cretaceous Kirkwood Formation, South Africa.

We name and describe a new iguanodontian dinosaur from the Early Creteceous Kirkwood Formation, Eastern Cape Province, South Africa. This dinosaur is one of only two ornithopod dinosaurs known from the Cretaceous of southern Africa, and is unique in being represented primarily by hatchling to young juvenile individuals as demonstrated by bone histological analysis. All of the juvenile material of this new taxon comes from a single, laterally-restricted bonebed and specimens were primarily recovered as partial to complete single elements, although rare articulated materials and one partial skeleton were found. Sedimentology of the bonebed suggests that this horizon heralds a change in environment upsection to a drier and more seasonal climate. This accumulation of bones is interpreted as seasonal mortality from a nesting site or nesting grounds and may be linked to this environmental shift.

Description of the first definitive Corythosaurus (Dinosauria, Hadrosauridae) specimens from the Judith River Formation in Montana, USA and their paleobiogeographical significance.

Despite the long history of research in the late Campanian Judith River Formation in northern Montana, most of the vertebrate fossils are represented by fragmentary remains, making precise taxonomic identifications difficult. Contrary to this, the partially contemporaneous Dinosaur Park Formation, Alberta, Canada is known for its tremendous fossil preservation, permitting rigorous studies of dinosaur diversity, evolution, and biostratigraphy. Hadrosaurids comprise one of the most abundant dinosaur clades in the Dinosaur Park Formation, but taxonomic affinities of hadrosaurid specimens remain poorly understood in the Judith River Formation. Corythosaurus is the most common hadrosaurid in the Dinosaur Park Formation and, to date, has been restricted to this formation. This study reports the first definitive Corythosaurus specimens from the Judith River Formation, which were discovered on two private ranches in northern Montana. The attribution of the most complete skeleton to Corythosaurus is indicated by: wide crest-snout angle, presence of premaxilla-nasal fontanelle, dorsoventrally expanded nasal, laterally exposed ophthalmic canal of the laterosphenoid, and tall neural spines. A second specimen preserves a large ilium that can be positively identified as Corythosaurus based on its associated skull, which is now in private hands. The specimens were recovered from the Coal Ridge Member of the Judith River Formation, which is approximately time equivalent to the Dinosaur Park Formation. Thus, the discovery of Corythosaurus in the Judith River Formation extends the biogeographic range of this genus and establishes a framework for future interformational biostratigraphic studies of Late Cretaceous dinosaur faunas in North America.